Phenotypic plasticity of sun and shade ecotypes of Stellaria longipes in response to light quality signaling: Cytokinins

From two distinct ecotypes of Stellaria longipes, one genotype was chosen from each of two very different locations, an alpine (sun) and a prairie (shade) habitat. Plants were clonally propagated and grown in controlled environment chambers under low and moderate red to far-red (R/FR) ratios. The prairie ecotype plants exhibited increased stem elongation, leaf expansion and flowering (6-fold) in response to a low R/FR ratio, relative to plants grown under the moderate R/FR ratio. In contrast, plants of the alpine ecotype showed no increased growth in response to a low R/FR ratio and their flowering was reduced, all relative to the plants grown under the moderate R/FR ratio. These different phenotypic responses to the reduction in R/FR ratio were associated with very different profiles and concentrations of endogenous cytokinins (CKs) assessed in growing tissues of the upper shoots. Specifically, increased total CKs were associated with the rapid growth of plants of the prairie ecotype under a low R/FR ratio. In particular, concentrations of bioactive trans-zeatin and dihydrozeatin, were increased during the period of most rapid shoot growth by 2- to 4- fold for these prairie ecotype plants grown under the low R/FR ratio treatment. In contrast, changes in CK levels for the alpine ecotype plants grown under low R/FR ratios were muted. Of especial interest, plants of the alpine ecotype had a predominance of cis-pathway CKs, whereas the low elevation, prairie ecotype plants accumulated predominantly trans-pathway CKs. Speculatively, the pattern emphasizing trans-pathway CKs may be explained by increased LONELY GUY enzyme activity. This enzyme converts and activates nucleotide CKs to free base CKs (bypassing riboside CKs). It could thus explain, in part, the prairie ecotype's ability to respond to shade light with such a high degree of plasticity if one assumes that high trans-CKs levels are causal for the increased shoot growth seen under a low R/FR ratio.     

Phenotypic plasticity in response to mechanical stress: hydrodynamic performance and fitness of four aquatic plant species

Plastic responses of plants exposed to mechanical stress can lead to modified, performance-enhancing, morphologies, sometimes accompanied by costs to reproduction. The capacity to present short-term plastic responses to current stress, the resulting performance (expected lower mechanical forces), and the costs of such responses to reproduction were tested for four aquatic plant species. Two ramets of the same genet were submitted to running vs standing water treatment. Traits describing the morphology, hydrodynamic performance and reproduction (sexual and vegetative) were measured. For one species, plastic responses led to reduced hydrodynamic forces, without apparent costs to reproduction, indicating that the plastic response could be beneficial for plant maintenance in stressful habitats. For two species, plastic responses were not associated with variations in performance and reproduction, possibly because of the low hydrodynamic forces experienced, even for morphologies produced under standing conditions. For one species, plastic responses were associated with a sharp decrease in sexual reproduction, without variations in performance, revealing the negative impact of currents over a short time scale. Species maintenance is linked to the capacity of individuals to tolerate mechanical forces. The contrasting responses to currents may be a key element for predicting community dynamics.     

Involvement of gibberellins in the stem elongation of sun and shade ecotypes of Stellaria longipes that is induced by low light irradiance

Plants from two ecotypes of Stellaria longipes, alpine (an open, sunny habitat) and prairie (where adjacent plants provide a shaded habitat), were grown under normal and reduced levels of photosynthetically active radiation (PAR). Growth under low PAR is significantly promoted in both ecotypes. When quantified by the stable isotope dilution method, endogenous gibberellins (GAs) (GA1, GA8, GA20, GA19) were significantly elevated under low PAR in both 'sun' and 'shade' ecotypes, as was GA53 in the shade ecotype. Changes in endogenous GA1 levels were significantly correlated with stem growth during a 28 d growth cycle and with relative growth rate (RGR) for height under low PAR for both ecotypes. Interestingly, under low irradiance PAR, changes (both increases and decreases) in GA8, the 2beta-hydroxylated 'inactive' catabolite of GA1, closely parallel bidaily stem growth changes for both ecotypes. Because the significantly greater stem elongation of both ecotypes in response to low irradiance PAR is associated with significant increases in the endogenous levels of five GAs (GA53, GA19, GA1, GA8) in the early 13-hydroxylation GA biosynthesis pathway (measured at days 7,14 and 21), we conclude that the low irradiance PAR has very likely induced an overall increase in GA biosynthesis.     

Phenotypic plasticity of internode elongation stimulated by deep-seeding and ethylene in wheat seedlings

Deep-seeding and ethylene were found to stimulate extension growth of the first internode of intact wheat (Triticum aestivum L.) seedlings in darkness. Seedlings of Hon Mang Mai emerged from much deeper in the soil than the seedlings of the other varieties used and their first internodes elongated to a much greater extent in response to ethylene. Carbon dioxide slowed elongation of the first internode and inhibited ethylene action. Elongation of the first internode due to deep-seeding and ethylene treatment showed high heritabilities, suggesting a genetic basis underlying those traits.     

Evolution of phenotypic plasticity: patterns of plasticity and the emergence of ecotypes

Phenotypic plasticity itself evolves, as does any other quantitative trait. A very different question is whether phenotypic plasticity causes evolution or is a major evolutionary mechanism. Existing models of the evolution of phenotypic plasticity cover many of the proposals in the literature about the role of phenotypic plasticity in evolution. I will extend existing models to cover adaptation to a novel environment, the appearance of ecotypes and possible covariation between phenotypic plasticity and mean trait value of ecotypes. Genetic assimilation does not sufficiently explain details of observed patterns. Phenotypic plasticity as a major mechanism for evolution--such as, invading new niches, speciation or macroevolution--has, at present, neither empirical nor model support.     

Growth and ethylene evolution by shade and sun ecotypes of Stellaria longipes in response to varied light quality and irradiance

Plants growing in the shade receive both low light irradiance and light enriched in far red (FR) (i.e., light with a low red (R) to FR ratio). In an attempt to uncouple the R/FR ratio effects from light irradiance effects, we utilized Stellaria longipes because this species has two distinct natural population ecotypes, alpine (dwarf) and prairie (tall). The alpine population occupies the open, sun habitat. By contrast, the prairie population grows in the shade of other plants. Both 'sun' and 'shade' ecotypes responded with increased stem elongation responses under low irradiance, relative to growth under 'normal' irradiance, and this increased growth was proportionally similar. However, only the shade ecotype had increased shoot elongation in response to a low R/FR ratio. By contrast, the sun ecotype showed increased stem elongation in response to increasing R/FR ratio. Varying the R/FR ratios had no significant effect on ethylene evolution in either sun or shade ecotype. Under low irradiance, only the sun ecotype showed a significantly changed (decreased) ethylene evolution. We conclude that R/FR ratio and irradiance both regulate growth, and that irradiance can also influence ethylene evolution of the sun ecotype. By contrast, R/FR ratio and irradiance, while having profound influences on growth of the shade ecotype, do not appear to regulate these growth changes via effects on ethylene production.     

喜旱莲子草对模拟全天增温的可塑性: 引入地和原产地种群的比较

植物可以通过关键功能性状的表型可塑性来适应气候变暖背景下环境温度的增加。表型可塑性增强进化假说(evolution of increased phenotypic plasticity hypothesis)认为外来植物在引入地进化出了更强的表型可塑性。以往对该假说的验证多集中于外来植物对光照、水分、养分、邻体以及天敌等的可塑性进化, 而对增温条件下植物生长和功能性状可塑性进化的研究相对较少。仅有的几项研究多集中在温带地区, 且多集中于研究植物生长相关的性状, 而对植物的抗性和草食作用对增温的响应的关注相对较少。本研究采用同质园实验比较了喜旱莲子草(Alternanthera philoxeroides)引入地(中国)和原产地(阿根廷)各8个种群的生物量、功能性状和草食作用在热带地区(广州市增城区)对模拟全天增温2℃的响应差异。结果表明: (1)模拟全天增温显著降低了喜旱莲子草总生物量(-7.8%)、贮藏根生物量(-12.8%)、分枝强度(-11.6%)和茎端取食率(-34.4%)。(2)模拟全天增温造成的引入地种群总生物量降低幅度大于原产地种群; 模拟全天增温使引入地种群的比茎长和茎端取食率降低, 而原产地种群则相反。(3)无论是否模拟全天增温, 引入地种群的贮藏根生物量(+31.5%)、分枝强度(+38.5%)、比茎长(+30.2%)、根冠比(+24.5%)和比叶面积(+20.0%)均高于原产地种群, 茎端取食率则低于原产地种群(-35.8%)。这些结果表明, 热带地区全天增温2℃对喜旱莲子草是一种胁迫; 引入地种群的生物量对模拟全天增温2℃的响应更强, 而其株形相关性状(比茎长)和草食作用(茎端取食率)对模拟全天增温的可塑性方向与原产地种群相反。由于引入地种群在热带地区模拟全天增温条件下生物量的下降和草食作用的增加明显高于原产地种群, 因此在未来全球气候变暖的背景下, 热带地区温度升高可能不利于喜旱莲子草种群多度的增加。     

松嫩平原两个生态型羊草实验种群对盐碱胁迫的生理响应

采用盆栽实验,对灰绿型和黄绿型两个生态型羊草(Leymus chinensis)在幼苗期分别用不同强度的NaCl、Na2CO3和混合盐碱胁迫,测定其叶片叶绿素含量、电解质外渗率和植株脯氨酸、Na／K等生理指标。结果表明,在3种盐碱胁迫条件下,灰绿型和黄绿型羊草的叶片叶绿素含量、电解质外渗率和植株脯氨酸、Na／K均与胁强之间呈直线y=a+bx变化,达到了显著或极显著水平,两个实验种群均具有耐盐碱能力,叶片叶绿素含量、电解质外渗率和植株脯氨酸、Na／K均有显著或极显著差异,它们在耐盐碱生理特性上已产生了分化,其中,灰绿型的耐盐碱能力强于黄绿型,本文为羊草生态型分化主导生态因子是土壤盐碱所致提供了的生理学证据。     

Phenotypic plasticity and integration in response to flooded conditions in natural accessions of Arabidopsis thaliana (L.) Heynh (Brassicaceae)

Flood response is a crucial component of the life strategy of many plants, but it is seldom studied in non-flooded tolerant species, even though they may be subjected to stressful environmental conditions. Phenotypic plasticity in reaction to environmental stress affects the whole plant phenotype and can alter the character correlations that constitute the phenotypic architecture of the individual, yet few studies have investigated the lability of phenotypic integration to water regime. Moreover, little has been done to date to quantify the sort of selective pressures that different components of a plant's phenotype may be experiencing under contrasting water regimes. Genetic differentiation and phenotypic plasticity at the single-trait and multivariate levels were investigated in 47 accessions of the weedy plant Arabidopsis thaliana, and the relationship of plastic characters to reproductive fitness was quantified. Results indicate that these plants tend to be highly genetically differentiated for all traits, in agreement with predictions made on the basis of environmental variation and mating system. Varied patterns of apparent selection under flooded and non-flooded conditions were also uncovered, suggesting trade-offs in allocation between roots and above-ground biomass, as well as between leaves and reproductive structures. While the major components of the plants' multivariate phenotypic architecture were not significantly affected by environmental changes, many of the details were different under flooded and non-flooded conditions.     

喜旱莲子草对同基因型邻体根系的表型可塑性: 入侵地和原产地的比较

植物对邻体根系的表型可塑性是指与无邻体对照相比, 即使个体平均可获取土壤资源相同, 在有邻体根系存在时植物也会改变根系生物量分配, 并影响其他功能性状和适合度。表型可塑性进化假说(evolution of plasticity hypothesis)认为外来植物在入侵地进化出了更强的表型可塑性。对该假说的验证多集中于外来植物对光照、水分、养分以及天敌等的可塑性进化, 但对邻体根系的可塑性在入侵植物中是否发生进化尚未见报道。我们采用同质园实验比较了喜旱莲子草(Alternanthera philoxeroides)入侵地(美国)和原产地(阿根廷)各5个基因型的适合度与功能性状对同基因型邻体根系的可塑性。结果表明: 喜旱莲子草的根冠比(P = 0.088)和比叶面积(P = 0.007)对同基因型邻体根系的可塑性在入侵地和原产地基因型间存在差异: 入侵地基因型在有邻体根系时根冠比和比叶面积增加, 而原产地基因型则相反。但是, 总生物量、贮藏根生物量、比茎长和分枝强度对邻体根系的可塑性在入侵地和原产地间没有显著差异。此外, 与分隔邻体根系相比, 同基因型邻体根系存在时总生物量(+9.9%)和贮藏根生物量(+13.9%)显著增加, 比茎长(-9.5%)显著降低。最后, 与原产地基因型相比, 总体上入侵地基因型的总生物量(+62.0%)和贮藏根生物量(+58.9%)增加, 比茎长(-28.5%)和分枝强度(-42.8%)降低。这些结果表明喜旱莲子草入侵地基因型与资源利用相关功能性状(如根冠比和比叶面积)对邻体根系的可塑性方向与原产地基因型相反; 但适合度和株型相关性状(如比茎长和分枝强度)对同基因型邻体根系的可塑性与原产地没有差异。     

喜旱莲子草在青藏高原对模拟增温的可塑性: 引入地和原产地种群的比较

气候变暖背景下植物可通过关键性状的表型可塑性来适应环境温度的增加。表型可塑性增强进化假说预测定植到新环境中的入侵植物种群具有演化出更强表型可塑性的潜力。此前对可塑性进化的研究涵盖了外来植物性状对水分条件、光照变化、土壤养分、邻体根系以及天敌防御等的响应, 而较少有研究关注增温条件下植物重要性状的可塑性进化。已有的部分研究多集中在温带和热带地区, 而较少关注入侵植物在高寒地区对增温的响应; 且研究多集中在植物生长相关性状, 较少关注功能性状和防御性状。本研究采用同质园实验比较了喜旱莲子草6个引入地(中国)种群和6个原产地(阿根廷)种群, 在西藏拉萨模拟全天增温2℃处理下的适合度性状、功能性状和防御性状的响应差异。结果表明: (1)高寒地区模拟全天增温显著提高了喜旱莲子草总生物量(+36.4%)、地上生物量(+34.5%)、贮藏根生物量(+51.4%)和毛根生物量(+33.6%), 降低了分枝强度(-19.8%)和比茎长(-30.2%); (2)模拟全天增温使引入地种群的比叶面积和黄酮含量增加, 而原产地种群则相反。这些结果表明高寒地区全天增温2℃对喜旱莲子草可能是一种有利条件。引入地种群的适合度性状对模拟全天增温2℃的响应比原产地种群更强, 而其光能利用相关性状和防御性状的响应可能提升了其在高寒地区的适合度。因此, 在未来全球气候变暖的背景下, 高寒地区温度升高可能更有利于喜旱莲子草引入地种群的定植和扩散。     

Developmental and genotypic differences in the response of pea stem segments to auxin

The objective of this investigation was to examine the response to exogenous auxin (indole-3-acetic acid; IAA)of stem segments at two developmental stages. The standard auxin response of excised stem segments and intact plants consists of an initial growth response and a prolonged growth response. We found that this biphasic response does not occur in internodes at very early stages. Stem segments of light grown pea of various genotypes were cut when the fourth internode was at 6–13% of full expansion (early-expansion) or at 18–25% of full expansion (mid-expansion). Length measurements of excised segments were made after 48 hours of incubation on buffer with or without auxin. An angular position transducer linked to a computerized data collection system provided high-resolution measurement of growth of stacks of segments incubated in buffer over 20 hours. Early-expansion segments of all genotypes deviated from the standard auxin response, while mid-expansion segments responded in a manner consistent with previous reports. Early-expansion segments of tall, light-grown plants were unique in showing an auxin-induced inhibition of growth. The auxin-induced inhibition correlated with high endogenous auxin content, as determined by HPLC and GC/MS, across genotypes and between early-expansion and mid-expansion segments of tall plants. Measurement of ethylene evolved from stem segments in response to auxin, and treatment of segments with the ethylene action inhibitor, norbornadiene, showed the inhibition to be mediated in part by heightened ethylene sensitivity. Growth of early-expansion segments of dwarf and severe dwarf plants was stimulated by exogenous auxin, but the growth rate increase was delayed compared to that in mid-expansion segments. This is the first time that such a growth response, termed the delayed growth response has been emonstrated. It is concluded that developmental stage and endogenous hormone content affect tissue response to exogenous auxin.     

Phenotypic plasticity of Capsella bursa-pastoris (Brassicaceae) and its effect on fitness in response to temperature and soil moisture

Capsella bursa-pastoris is one of the most common plants on earth. Although phenotypic plasticity of ecologically important traits possibly contributes to its wide geographic range, little is known about the plasticity of C. bursa-pastoris and its effects on its fitness. In a laboratory, we assessed the phenotypic plasticity in response to two representative climatic conditions: temperature and soil moisture. In addition, we quantitatively evaluated the relationship between phenotypic plasticity and fruit production. Most measured morphological and physiological traits exhibited plastic responses to temperature and fitness based on fruit production was maintained across temperatures. In contrast, no plasticity to soil moisture was detected, and plants produced fewer fruits in dry soil. Selection analysis revealed that the plasticity of the flowering time and the water-use efficiency had positive effects on fruit production over the tested temperature regime. These experimental results suggest that phenotypic plasticity probably enables C. bursa-pastoris to cope with heterogeneous temperature environments and thereby probably contributes to its wide geographic range.     

Sensitivity to ethylene: the key factor in submergence-induced shoot elongation of Rumex

Rosettes of flooding-resistant Rumex palustris plants show a submergence-induced stimulation of elongation, which is confined to the petioles of young leaves. This response increases the probability of survival. It is induced by ethylene that accumulates in submerged tissues. Flooding-intolerant Rumex acetosella plants do not show this response. We investigated whether differences in shoot elongation between the species, between old and young leaves and between the petiole and leaf blade of a R. palustris plant result from differences in internal ethylene concentration or in sensitivity to the gas. Concentrations of free and conjugated ACC in petioles and leaf blades of R. palustris indicated that ethylene is synthesized throughout the submerged shoot, although production rates varied locally. Nevertheless, no differences in ethylene concentration were found between submerged leaves of various ages. In contrast, dose-response curves showed that only elongation of young petioles of R. palustris was sensitive to ethylene. In R. acetosella, elongation of all leaves was insensitive to ethylene. We conclude that variation in ethylene sensitivity rather than content explains the differences in submergence-induced shoot elongation between the two Rumex species and between leaves of R. palustris.     

Phenotypic plasticity and the possible role of genetic assimilation: Hypoxia-induced trade-offs in the morphological traits of an African cichlid

In this study we investigate the possible role of phenotypic plasticity and genetic assimilation in the process of adaptation and evolutionary change in the cichlid Pseudocrenilabrus multicolor victoriae . In the field we compared a population of a stable hypoxic habitat with one of a stable well-oxygenated habitat. In the laboratory, we compared individuals from the same mother raised under hypoxic or well-oxygenated conditions to examine phenotypic plasticity. Morphological parameters of three categories were measured: (a) the gill apparatus, (b) the surrounding structural elements, and (c) the outer shape of the fish. Swamp-dwelling fish had a 29% greater total gill surface area than fish from the well-oxygenated habitat due to their larger gill filament length and greater lamellar area. In the plasticity experiment, total gill surface area was 18% greater in the hypoxia group due to a larger number of longer filaments. Surrounding elements and outer shape also differed between the field populations and between fish grown under hypoxic and well-oxygenated conditions, but there was disparity between the field results and the plasticity experiment. The disparity between field and experimental fish may be due to: (a) differences in selection pressures between populations, (b) different constraints for genetic and plasticity changes, or (c) selection against plastic responses to hypoxia. Our results suggest that both (a) and (c) are involved.     

Morphological plasticity in the kelp Nereocystis luetkeana (Phaeophyceae) is sensitive to the magnitude,direction, and location of mechanical loading

Nereocystis luetkeana is a canopy-forming kelp that exhibits morphological plasticity across hydrodynamic gradients, producing broad, undulate blades in slow flow and narrow, flattened blades in fast flow, enabling thalli to reduce drag while optimizing photosynthesis. While the functional significance of this phenomenon has been well studied, the developmental and physiological mechanisms that facilitate the plasticity remain poorly understood. In this study, we conducted three experiments to characterize how the (1) magnitude, (2) direction, and (3) location of plasticity-inducing mechanical stimuli affect the morphology of Nereocystis blades. We found that applying a gradient of tensile force caused blades to grow progressively longer, narrower, less ruffled, and heavier in a linear fashion, suggesting that Nereocystis is equally well adapted for all conditions within its hydrodynamic niche. We also found that applying tension transversely across blades caused the growth response to rotate 90°, indicating that there is no substantial separation between the sites of stimulus perception and response and suggesting that a long-distance signaling mechanism, such as a hormone, is unlikely to mediate this phenomenon. Meristoderm cells showed morphological changes that paralleled those of their respective blades in this experiment, implying that tissue-level morphology is influenced by cell growth. Finally, we found that plasticity was only induced when tension was applied directly to the growing tissue, reinforcing that long-distance signaling is probably not involved and possibly indicating that the mechanism on display generally requires an intercalary meristem to facilitate mechanoperception.     

Phenotypic plasticity and the evolution of trade-offs: the quantitative genetics of resource allocation in the wing dimorphic cricket, Gryllus firmus

In the wing dimorphic sand cricket, Gryllus firmus, there is a pronounced trade-off between flight capability and fecundity. This trade-off is found both between morphs and within the macropterous morph, in which fecundity is negatively correlated with the mass of the principle flight muscles, the dorso-longitudinal muscles (DLM). In this paper, we examine how this trade-off is affected by a reduction in food and its genetic basis. We find that the relative fitness of the two wing morphs is not changed although both fecundity and DLM mass are decreased. A quantitative genetic analysis shows that the trade-off function is genetically variable but that most of the variation occurs in the intercept rather than the slope of the function. Analysis further indicates a very high genetic correlation between environments (food ration) supporting the hypothesis of a strong functional constraint between reproduction and flight capability.     

Evolution in stressful environments II: adaptive value and costs of plasticity in response to low light in Sinapis arvensis

Plants possess a remarkable capacity to alter their phenotype in response to the highly heterogeneous light conditions they commonly encounter in natural environments. In the present study with the weedy annual plant Sinapis arvensis, we (a) tested for the adaptive value of phenotypic plasticity in morphological and life history traits in response to low light and (b) explored possible fitness costs of plasticity. Replicates of 31 half-sib families were grown individually in the greenhouse under full light and under low light (40% of ambient) imposed by neutral shade cloth. Low light resulted in a large increase in hypocotyl length and specific leaf area (SLA), a reduction in juvenile biomass and a delayed onset of flowering. Phenotypic selection analysis within each light environment revealed that selection favoured large SLA under low light, but not under high light, suggesting that the observed increase in SLA was adaptive. In contrast, plasticity in the other traits measured was maladaptive (i.e. in the opposite direction to that favoured by selection in the low light environment). We detected significant additive genetic variance in plasticity in most phenotypic traits and in fitness (number of seeds). Using genotypic selection gradient analysis, we found that families with high plasticity in SLA had a lower fitness than families with low plasticity, when the effect of SLA on fitness was statistically kept constant. This indicates that plasticity in SLA incurred a direct fitness cost. However, a cost of plasticity was only expressed under low light, but not under high light. Thus, models on the evolution of phenotypic plasticity will need to incorporate plasticity costs that vary in magnitude depending on environmental conditions.     

Phenotypic plasticity to light intensity in Arabidopsis thaliana: invariance of reaction norms and phenotypic integration

Phenotypic plasticity (the pattern of response of organisms to changes in environmental conditions) and phenotypic integration (the pattern of character correlations) are important components of our understanding of the evolution of complex phenotypes. Most studies published so far in this area have been conducted within populations with the express aim of predicting future response to evolutionary forces. However, among-population differentiation for plasticity and trait correlations are important indicators of recent past events that have shaped the currently observable phenotypes. We investigated variation in the reaction norms of several traits in a large number of accessions of Arabidopsis thaliana exposed to different levels of light quantity as well as the environmental lability of the corresponding across-population character variance–covariance matrix. Our results show that there is an astounding degree of inter-population variation for character means and very little variation for plasticity, in agreement with the idea that A. thaliana is a light-specialist often occurring in open, disturbed habitats. However, this plant also shows patterns of plasticity that are predicted to be adaptive based on functional ecological considerations, such as an increase in either specific leaf area or leaf number (but not both) under low light. We also demonstrate that the set of character correlations in A. thaliana is extremely stable to changes in light availability, contrary to previous findings in the same species when different environmental factors were considered. Several processes that might have been responsible for the observed patterns are discussed as a prelude to follow-up research on these problems.     

The role of leaf lobation in elongation responses to shade in the rosette-forming forb Serratula tinctoria (Asteraceae)

BACKGROUND AND AIMS: Lobed leaves are considered selectively advantageous in conditions of high irradiance. However, most studies have involved woody species, with only a few considering the role of leaf lobation in herbaceous rosette species. In this study, it is hypothesized that, in addition to its adaptive value in high light, leaf lobation may add to the function of petioles as vertical spacers in herbaceous species in conditions of strong competition for light. METHODS: To test this hypothesis, leaf development was examined under seasonally changing natural light conditions and a field experiment was conducted in which light climate was manipulated in a wooded meadow population of Serratula tinctoria. KEY RESULTS: No changes in leaf lobation were observed in response to experimental shading or different natural light conditions. However, in tall herbaceous vegetation, plants with highly lobed leaves achieved significantly greater vertical elongation than plants with less-lobed leaves. In contrast to herbaceous shade, tree shade had no effect on leaf elongation, suggesting differential responsiveness to competition from neighbouring herbs versus overhead shade. In shading treatments, imposed shade could only be responded to by the elongation of leaves that were produced late in development. CONCLUSIONS: The results show that extensive leaf lobation can enable greater leaf elongation in response to shade from surrounding herbaceous vegetation. The different morphological responses displayed by Serratula tinctoria to different types of shade demonstrate the importance of critically assessing experimental designs when investigating phenotypic plasticity in response to shade.